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Responses of the Fish Blennius Pholis to Fluctuating Salinities

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Responses of the Fish Blennius Pholis to Fluctuating Salinities Vol. 1, 101-107, 1979 MARINE ECOLOGY - PROGRESS SERIES Published September 30 Mar. Ecol. hog. Ser. Responses of the Fish Blennius pholis to Fluctuating Salinities J. Davenport and 0. Vahl* N.E.R.C. Unit of Marine Invertebrate Biology, Marine Science Laboratories, Menai Bridge, Gwynedd LL59 5EH. United Kingdom ABSTRACT: Blennius pholis (L.) were exposed to fluctuating salinity regimes of near tidal periodicity and their blood osmolality, oxygen consumption, heart rate and opercular beat monitored. Despite salinity fluctuations from 34-0-34 '/m S, the blood osmolality remained constant. Significant increases in oxygen consumption were observed at low salin~tylevels, but these may simply reflect changes in physical activity and the greater availability of oxygen at low salinities. Salinity effects upon heart rate/ opercular beat were weak or negligible. INTRODUCTION imposed upon animals by the salinity'of their environ- ment. A number of invertebrates can effectively coun- The blenny Blennius pholis (L.) is a common inhabi- teract salinity stress during periods of each tidal cycle tant of rocky shores around Anglesey, U.K., particu- in littoral and estuarine areas, and thereby avoid or larly on the west coast of the island. It lives in rocky reduce damage by low environmental salinity levels. pools beneath stones, buried in muddy gravel and in Such compensations have been considered by Kinne rock crevices from just above the low water springtide (1967, p. 526) under the headings of escape, reduction level to the upper shore. Although it is a non-grega- of contact, regulation and acclimation. In several cirri- rious fish (Gibson, 1967a, 1968), its defended home pede, bivalve and polychaete species, compensation ranges are small, and thus local abundances may be for salinity stress has been studied by Davenport quite high. Blennies are particularly numerous in nar- (1976a), Davenport (1977) and Shumway and Daven- row rock gullies running at right angles to the coast, port (1977).As suggested by Kinne (1971b, p. 905) and and these are often fed by freshwater streams or see- Spaargaren (1974), invertebrates which cannot reduce page. Consequently the fish are exposed to considera- contact with adverse salinities tend to exhibit internal ble salinity fluctuations ranging from 33 to 34 O/OO S at osmolarity changes which are damped by comparison high tide, to freshwater at low tide. The individuals with those of the medium, and thus allow the animals living highest on the shore may be exposed to freshwa- to survive brief exposures to salinity levels which ter influence for several hours. would kill them if sustained for periods longer than The information available on the effects of salinity those prevailing in their natural environment. Such upon marine organisms in terms of their tolerance and damping has been confirmed by Tucker (1970) for the physiology has been reviewed extensively in 'Marine prosobranch Scutus breviculus, and for echinoderms Ecology', Volume I (Kinne, 1971a) and has received and bivalves by Stickle and Ahokas (1974) and Shum- detailed attention notably from Krogh (1939),Potts and way (1977). Data on fluctuating salinity effects upon Parry (1964) and Kinne (1971b). However, almost all of organisms have been obtained with the aid of an appa- the studies conducted thus far have involved labora- ratus which can mimic environmental salinity fluctua- tory experiments at constant salinities. tions: the most convenient and accurate apparatus is Steady-state salinity experiments are open to criti- that described by Davenport et al. (1975). cism on several grounds. Primarily they tend to give a Most of the information available thus far has been false impression of the possible limits to distribution obtained in invertebrates, largely for osmoconformers rather than osmoregulators. In teleost fishes the situa- On leave from Institute of Biology and Geology, University tion is different. They have a low blood concentration of Tromso, Norway. maintained by an efficient osmoregulatory mechanism O by Inter-Research 102 Mar. Ecol. Prog. Ser., 1, 101-107, 1979 (e.g. Gilles, 1975) which, in euryhaline species, is rometric experiments described below, an abrupt responsible for the extrusion of salts at high salinity regime as described already, and a sinusoidal regime levels and for taking up salts at low seawater concen- of 12 hour wavelength fluctuating between full seawa- trations. The blenny appeared to be suitable for a first ter and fresh water. investigation with the fluctuating salinity apparatus, since this teleost is relatively small and its behaviour causes it to remain at positions on the shore which Heart/Opercular Beat Frequency Measurements expose it to local tidal salinity fluctuations. In the study reported here, blennies were exposed to Heart and opercular beat frequencies were both fluctuating salinity regimes of near-tidal periodicity, measured simultaneously in individual fish by the During this exposure their plasma osmolarity, oxygen impedance pneumograph technique (Trueman, 1967). consumption, heart rate and opercular beat frequen- Each fish was anaesthetised with MS 222 and a silver- cies were monitored. wire electrode sewn beneath the skin overlying each operculum. The two electrodes were connected, via fine polythene sheated copper wire to a Scientific MATERIALS AND METHODS Instruments' pneumograph coupler, itself connected to a twin channel George Washington 400 MD/2 chart Collection and Maintenance recorder. Another pair of silver electrode's were inser- ted through the body wall (either side of the heart); Blennies Blennius pholis (L.) were mainly collected they were anchored in place by sewing to the bases of from the intertidal zone of the rocky west coast of the pelvic fins. These electrodes were also connected Anglesey (U.K.) between Aberffraw and Rhosneigr. A to a pneumograph coupler and the twin channel recor- few individuals were obtained from the shore of the der. The fish was then allowed to recover for several Menai Strait beneath the northern end of Telford hours before use in experiments. To conserve chart Suspension Bridge. The fish were collected from pools, paper during long experiments the power supply of the crevices and muddy gravel beneath stones. Fish in chart recorder was controlled by a time clock which pools were caught by the use of the anaesthetic quinal- switched the recorder on for 2 min every hour. Results dine (Gibson 196713). The blennies ranged in wet were obtained for 3 fish (19.0-27.5 g) exposed for 24 h weight from 7.4-42.6 g. To obtain arrythmic fish, the to the abrupt salinity regime. blennies were held in the laboratory seawater system at 10 "C and under constant illumination (Gibson 1967a). They were fed flesh of Mytilus edulis regularly, Plasma Osmolality Determinations but starved for 2 days before use in respirometry expe- riments. All experiments were performed in constant Blood samples were obtained by sacrificing fish at light at 10 "C. appropriate intervals during the first 12 h of an abrupt salinity regime. The tail of each fish was cut off and blood allowed to drip into a 1.5 m1 Eppendorf plastic Salinity Regimes tube. After centrifuging the blood to remove cells, a 50 mm3 sample of plasma was pipetted into the test vessel The blennies were exposed to salinity fluctuations of Knauer Semi-micro osmometer and its osmolality produced by the apparatus described by Davenport et established. Enough blood could be obtained from a al. (1975).To determine the influence of these fluctua- single large fish for such a determination, but on tions upon heart/opercular beat frequencies and several occasions the blood of 2-3 smaller fish had to plasma osmolarity the fish were placed in 500 m1 be pooled. plastic boxes supplied with water at 250-300 m1 min-l. The contents of the boxes were aerated by diffuser Respirometry stones connected to the compressed air supply. In these experiments only one type of salinity regime was used, Oxygen consumption measurements were carried namely an abruptly fluctuating cycle in which the test out by the use of the apparatus shown in Figure 1. Air- fish received alternate 6 h exposures to full seawater saturated water was drawn by a peristaltic pump from (33.5 '/W S) and fresh water. No more gradual regimes a column supplied with water of fluctuating salinity by than this were employed because the monitored phy- the apparatus described by Davenport et al. (1975). It siological changes induced by the abrupt salinity fluc- tuations were either slight or negligible. Scientific Instrument Centre, 13 Derby Lane, Lverpool, Two types of salinity regime were used in the respi- U.K. Davenport and Vahl: Responses of Blennius pholis to fluctuating salinites column Fig. 1. Experimental set-up for measurement of oxygen consumption in various salinity regimes was then delivered to a 250 m1 glass experimental (5) The fish was removed from the respirometer and chamber fitted with a screw top plastic-coated metal weighed. lid, and magnetically stirred. From the experimental To avoid potential inaccuracies caused by oxygen- chamber the water flowed past a Radiometer E5046 electrode response drifts, the electrode was regularly oxygen electrode, connected via a Radiometer PHM 7 1 moved from its position in the experimental-chamber Mk I1 pH meter to a Smiths Servoscribe chart recorder outflow and placed for a few minutes in the surround- adjusted to 100 mV. The water then flowed through a ing temperature-controlled bath. Since this bath is platinum conductivity cell, connected via a Carwyn continually aerated, and at the same temperature as Instruments' Salinity Monitor to the same chart recor- the experimental chamber and aeration column, this der as the oxygen electrode. Thus a nearly simultane- procedure allows regular reassessment of the chart ous trace of the salinity and oxygen tension of the deflection corresponding to full air saturation. When water leaving the experimental chamber was avail- the flow rate of water through the respirometer is able.
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